Geol. Quart. 64 (4) Internet

Geo log i cal Quar terly, 2020, 64 (4): 876–897 DOI: http://dx.doi.org/10.7306/gq.1565 The his tory of bears (Ursidae, Carnivora, Mammalia) from Silesia (south ern Po land) and the neigh bour ing ar eas Adrian MARCISZAK1, * and Grzegorz LIPECKI2 1 Uni ver sity of Wroc³aw, De part ment of Palaeozoology, In sti tute of En vi ron men tal Bi ol ogy, Fac ulty of Bi o log i cal Sci ences, Sienkiewicza 21, 50-335 Wroc³aw, Po land 2 Pol ish Acad emy of Sciences, In sti tute of System at ics and Evo lu tion of An i mals, S³awkowska 17, 31-016 Kraków, Po land Marciszak, A., Lipecki, G., 2020. The history of bears (Ursidae, Carnivora, Mammalia) from Silesia (south ern Poland) and the neigh bour ing ar eas. Geo log i cal Quar terly, 64 (4): 876–897, doi: 10.7306/gq.1565 As so ci ate Ed i tor: Micha³ Zatoñ Re vi sion of the Silesian bear fauna, based on ma te rial from 152 sites, mainly cave and karstic lo cal i ties, and also ar chae o- log i cal and open-air sites, shows the presence of 13 forms and spe cies. These re cords en com pass the last 16.5 Ma and may be di vided into five main morphophyletic groups. The old est bears, rep resented by the genera Ballusia and Ursavus, are dated to 16.5–11 Ma, and belonged to the stem forms of the subfamily Ursinae. Af ter a break of 6 My, the earli est mem bers of the ge nus Ursus ap peared, and all known Plio cene bears in Silesia be long to the ge nus Ursus were identi fied as Ursus minimus. They repre sent one or two migra tion events. Ad di tion ally, between 3.6 and 3.2 Ma, a single oc cur rence of Agriotherium insigne was recorded from the Wê¿e 1 site. A few Early Pleis tocene bear records are repre sented by U. etruscus, which was a proba ble an ces tor of both arctoid and spelaeoid bear lin eages. The old est rep resen tative of Ursus ex gr. arctos known so far and assigned to U. a. suessenbornensis is known from one lat est Early Pleis tocene (1.2–0.9 Ma) lo cal ity, while other Pol ish re cords of this form re quire con fir ma tion. The first oc cur rence of U. deningeri, the old est taxon within the U. ex gr. deningeri–spelaeus lin eage, was recorded from ~700 ka de pos its in Silesia. Dur ing the pro nounced cold period of MIS 12, the Scan di na vian ice sheet cov ered al most the en tire mod ern terri tory of Po land, with the excep tion of the Sudetes and the Carpathians. The ac com pa nying dras tic fau nal turn over led to the for ma tion of the pan-Eurasian Mam moth Fauna at ~460 ka. At that time a char ac teris tic mem ber of this fauna, the steppe brown bear Ursus arctos priscus, a spe cific ecomorph adapted to live in open grass lands, ap peared in this re gion. It sur vived un til the be gin ning of MIS 1, when mod ern Ursus arctos arctos ap peared in Silesia and sur vived to the pres ent day. U. deningeri was the most com mon bear during the Mid dle Pleis to cene, while the first re cords of U. spelaeus spelaeus ap peared since MIS 7. The lat ter form was re placed by U. ingressus during the Late Pleisto cene (~110–100 ka). Spelaeoid bears to tally domi nated the cave as sem blage, and fi nally van ished be tween 27 and 24 ka. Key words: tax on omy, Mio cene, Plio cene, Pleis to cene, ursid lin eages. INTRODUCTION Bears spread all over the North ern Hemi sphere dur ing the Mio cene and gave rise to a con sid er able num ber of spe cies, from which a single one has sur vived to the pres ent in Po land. Finds of dif fer ent rep re sen ta tives of the fam ily Ursidae are The Silesian his tory of bears started with the ear li est Pol ish re - fre quently made in Po land, and their high est den sity has been cord of Ursavus elmensis from the Przeworno 2 site dated to re corded from the cave and karstic sites of Silesia and im me di - 16–14.5 Ma. This taxon is also gen er ally consid ered to be the ately neighbour ing ar eas. Palaeontologically this re gion is one ear li est un dis puted bear spe cies (Baryshnikov and Lavrov, of the most pro lific in Po land. Bear re cords from Silesia and 2015). The Mio cene ge nus Ursavus Schlosser, 1899 is re - nearby areas al low us to re con struct the re gional his tory of the garded as a di rect an ces tor of mod ern ur sine bears of the ge - fam ily over the last ~16 My. This timespan is, though, in com - nus Ursus Linnaeus, 1758. Bears in Silesia have been al most pletely cov ered, and there are some gaps, es pe cially in the mid - ex clu sively rep re sented by spe cies of Ursus over the last 4 My, dle Mio cene to Mid dle Pleis to cene in ter val. with the ex cep tion of a sin gle oc cur rence of Agriotherium that per sisted from the Mio cene (Stach, 1953, 1957; Wolsan, 1989). Ur sine bears have existed in Silesia con tin u ously since the Mid dle Plio cene, from ~4 Ma. Al though only sporadic in some in ter vals, bears be came one of the dom i nant mam mal groups * Cor re spond ing au thor, e-mail: [email protected] dur ing the Mid dle and Late Pleis to cene. This pa per pro vides a Re ceived: April 14, 2020; ac cepted: Sep tem ber 17, 2020; first gen eral over view of the his tory and taxon omy of ur sine bears in pub lished on line: November 12, 2020 Silesia and the near est neigh bour ing ar eas, as a crit i cal re view The his tory of bears (Ursidae, Carnivora, Mammalia) from Silesia (south ern Poland) and the neigh bouring ar eas 877 of the lit er a ture com ple mented by our own re search re sults. fam ily in the palaeofaunas grew sig nif i cantly, and from the Mid - Size and mor pho log i cal anal y ses are sup ple mented by ura - dle Mio cene bears be came im por tant mem bers of the nium-tho rium (UTh) and ac cel er a tor mass spec trom e try (AMS) palaeoguilds. By that time there had al ready been estab lished a dat ing meth ods as well as by DNA analy sis. model of bear evo lu tion, based on a large, solid an i mal, mostly om niv o rous or car niv orous, able to com pete with even the larg - est ac tive car ni vores like big cats. In the last 15 My on almost all MATERIAL AND METHODS con ti nents (ex pect Aus tra lia) a large bear, Agriotherium ear lier, Arctodus or Ursus later, which played an impor tant role as a The sub di vi sion of the fam ily Ursidae Fischer, 1814 into five scav en ger of and com pet i tor to other large car ni vores. subfamilies: Ursinae Fischer, 1814, Arctotheriinae F. Dur ing this evo lu tion, the size of mem bers of the Ursinae Ameghino, 1903, Ailuropodinae Grevé, 1892, Agriotheriinae subfamily also grew con sid er ably (Fig. 2). The ear li est Silesian Kretzoi, 1929, and Ursavinae Kretzoi, 1945 is made sensu bear, B. elmenensis, was the size of a lynx, while their di rect de - Wag ner (2010). Ac cord ing to this con cep tion, all Eur asian rep - scen dants of the ge nus Ursavus were al ready the size of a re sen ta tives of the Ursinae subfamily discussed in this pa per large wolf. Plio cene forms like U. minimus were com pa ra ble in may be assigned to the ge nus Ursus. The tax o nomic over view size with the modern Hi ma la yan black bear Ursus thibetanus was made ac cord ing to main groups of Eur asian bears sensu Cuvier, 1823. De spite its rel a tively large di men sions, A. in signe Mazza and Rustioni (1994): (1) minimus–thibetanus, (2) from the Wê¿e 1 site is a me dium-sized mem ber of the ge nus etruscus, (3) arctos, and (4) deningeri–spelaeus. The def i ni tion Agriotherium. The Early Pleis to cene U. etruscus reached the and sub di vi sions of the Qua ter nary Pe riod fol low Gibbard and size of a mod ern Cen tral Eu ro pean brown bear. Ursids reached Co hen (2008) and Gibbard and Head (2009a, b). The def i ni tion their maxi mum size in the Mid dle and es pe cially Late Pleis to - and sub di vi sions of mam mal zones and their cor re la tion with cene, where deningeroid and spelaeoid bears dom i nated in the chronostratigraphic scale as well as the MN-zones fol low caves and were the larg est Eur asian bears.

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